seedling ecology and · pdf file 10.3 understanding seed and seedling ecology as parts of a...

Seedling Ecology and Evolution

Editors

Mary Allessio Leek Emeritus Professor of Biology, Rider University, USA

V. Thomas Parker Professor of Biology, San Francisco State University, USA

Robert L. Simpson Professor of Biology and Environmental Science, University DJ Michigan - Dearborn, USA

~CAMBRIDGE V UNIVERSITY PRESS

Contents

Contributors

Foreword by Perer J. Grubb Preface Acknowlrdgments

Part I I Introductio n

Chapter I I Why seedlingsr Mary AUessio Leek, Roberr L Simpson. and V. Thomas Parker

1.1 Seedlings as part of a plant's life cycle 1.2 Vulnerabilities and bottlenecks 1.3 Making ie fil te rs, safe sires. and establish ment 1.4 Seedlings: a primer 1.5 What seedlings can tell us 1.6 The scope of Seedling Ecology and Evolution

Part 11 I Seedling dive rsity

Chapter 2 I Seedling natural history Mary Ai!essio Leek and Heather A. Outred

pagt xi xiii

,~I

xix

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3

6

7

8

10

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" 2.1 Introduction 17 2.2 The seedling stage and fate of seedlings 17 2.3 Seedling types 20 2.4 Seedling diversity - morphology 24 2.5 Seedling diversi ty - ecophysiology 31 2.6 Vivipary and seedling equ ivalents 33 2.7 Longevity 36 2.8 Dispersal 37 2.9 Environmental filters and safe siles 38 2.10 Summary 41 2.11 Acknowledgments 41

Appendixes 42

Chapter 3 I Specialized seedling strategies I: seedlings in stressful environments 56

josi M. Facdli 3.1 Introduction 3.2 Seedling establish ment ill dense shade 3.3 Effects of liner on seedling establishment 3.4 Seedling establishment in dry environments 3.5 Seedling establishment in saline envi ronments

" 60 " 6S 68

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,; CONTENTS

3.6 Seedling establishment in cold environments 70 3.7 Physical damage to seedlings 73 3.8 Interactive effects of stress agents and biological

interactions 75 3.9 Ovetview: adaptations of seedlings to stressful

environments 76 3.10 Acknowledgments " Chapter 4 1 Specialized seedling st rategies 11:

orchids, bromeliads, carnivorous plants, and parasites 79

Dermis F. Whigham, Mr!lssa K. McCormick, • and John P. O'Neill

4.1 Introduction 79 4.2 Epiphytic and terrestrial orchids 80 4.3 Bromeliads 85 4.4 Carnivorous plants 88 4.5 Parasitic plants 91 4.6 Summary and future needs " 4.7 Acknowledgments 100 Part III I Seedling morphology, evolution,

and physiology

Chapter 5 I Embryo morphology and seedling evolut ion 103 Karl J. Nilelas

5.1 Introduction 103 5.2 Embryophyte phylogeny 106 5.3 Embryo structure 108 5.4 Embryophyte embryogenesis 112 5.5 Phylogenetic patterns 122 5.6 Concluding remarks 128

Chapter 6 I Regeneration ecology of early angiosperm seeds and seedlings: integrating inferences from extant basal lineages and fossils 130

Taylor S. Fe/Id 6.1 Introduction 130 6.2 Previous views of ancestral angiosperm ecology and

seed/seedli ng morphology 132 6.3 The phylogenetic revolution :.inferences on early

angiosperm regeneration ecology from extant basal angiosperms 13'

6.4 FUnctional biology of basal angiosperm seeds 136 6.5 Functional biology of basal angiosperm seedlings 1

Chapter 7 I Physiological and morphological changes during early seedling growth: roles of phytohormones

Elizabeth J. Farnsworth 7.1 Introduction: phytohormones, molecular biology,

150

and the ~real world M of early seedling ecology 150 7.2 Seedling responses to light 156 7.3 Seedling responses to temperature 159 7.4 Seedling responses to water 161 7.5 Seedling responses to nutrients 163 7.6 Insights and common themes 166 7.7 Summary 170 7.B Acknowledgments 171

Chapter 8 I Seedling ecophysiology: strategies toward achievement of positive net carbon balance 172

Kaom Kitajima and Jonathan A. Myers 8.1 Introduction 172 8.2 Seed reserve utilization 173 8.3 Ontogenetic trajectories of seedling carbon balance 175 8.4 Species differences in inherent relative growth rate

(RGR) 177 8.5 Opportunistic versus conservative strategies 178 B.6 Carbohydrate reserves 182 8.7 Phenotypic plasticity 185 8.8 Concluding remarks 187 B.9 Acknowledgments 188

Chapter 9 I The role of symbioses in seedling establishment and survival

Thomas R. Horton and Marcr1 G. A. \lan der Heijden

189

9.1 Introduction 189 9.2 Ectomycorrhizal fungi and seedling establishment 199 9.3 Arbuscular mycorrhizal fungi and seedling

establishment 207 9.4 Other plant symbionts and seedling establishment 209 9.5 Conclusions 212 9.6 Acknowledgments 213

Part IV I Life history implications

Chapter 10 I The seedling as part of a plant's life history strategy

Ange!a T. Mo!es and Michel!e R. ieishman 10.1 Introduction

217

CONTENTS vii

viii CONTENTS

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10.2 The trade-off be£Ween offspring production and seedling survival 218

10.3 Understanding seed and seedling ecology as parts of a plant's life history stra tegy 227

lOA Correlations between seed and seedling strategy and other aspects of plant ecology 229

10.5 Seed and seedling strategIes within species 232 10.6 Implications of a holistic understanding of plant life

history strategies 235 10.7 Conclusions 237

Chapte r 1 I I Seedling recruitment and population ecology

Ove Etiksson and Johan fhrlen 11.1 Introduction 11.2 TIu~ causes of seedling mortality 11.3 Recruitment limitation 11.4 Seedling recruitment and population dynamics 11.5 Genetic structure and selection in seedling

populations 11.6 Concluding remarks 11.7 Acknowledgments

Chapte r 12 I Seedling communities Jon F.. Keeler and PhillipJ. van Mantgt'lTl

12.\ Introduction 12.2 Internal drivers 12.3 External drivers affecting seedling communities 12.4 Seedling communil}' assembly rules 12.5 Conclusions

Chapte r 13 I Spatial variation in seedling emergence and establishment - functional groups among and within habitats~

Johannes Kollmann 13.1 Introduction 13.2 Description of the seedling stage 13.3 Definition of spatial scales 1304 Microhabitat effects on seedling dynamics 13.5 Habitat effects on seedling dynamics 13.6 Landscape effects on seedling dynamics 13.7 Region effects on seedling dynamics 13.8 Biome effects on seedling dynamics: seed site and

seedling survival 13.9 Synthesis 13.10 Acknowledgments

239

239

241

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Part V I Applications

Chapter 14 I Does seedling ecology matter for biological invasions!

Laura A. Hyatt 14.1 Introduction 14.2 Invasive seedlings 14.3 Invasive effccts on native seedlings 14.4 Conclusions 14.5 Acknowledgments

Chapter IS I The role of seedlings in the dynamics of dryland ecosystems - their response to and involvement in dryland heterogeneity,

295 296

303

305

306

degradation, and restoration 307

Bertrand Boeken 15.1 Introduction 307 15.2 Importance of the seedling stage 308 15.3 Seedlings and spatial heterogeneity of drylands 312 15.4 Seedlings and dryland system degradation 322 15.5 Conclusions 330

Chapter 16 I Anthropogenic disturbance in tropical forests: toward a functional understandi ng of seedling responses 332

james W Da!!ing and David F. R. r. Rurslem 16.1 Introduction 332 16.2 Significance of the seedling stage for forest

management 16.3 Effe

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17.5 Mimicking the effects of disturbances in restoration 368 17.6 Conclusions 369

Pa,.t VI I Synthesis

Chapter 18 I The seedling in an ecological and evolutionary context

V Thomas ParkIT, Robert L. Simpson, and Mary Al!essio Leck

373

18.1 Introduction 373 18.2 Dispersal. seed bank dynamics, and seedling banks 374 18.3 Dynamics of individual seedlings 376 18.4 Set'dlings in heterogeneous environments 378 18.5 Alternative strategies 381 18.6 Conclusions 385

References Index

391

501

Index

ARA. see abscisic acid Abies. 36,119,265

A aloo. ]52

A amabUis. 202

A conrol'or, 257-9, 268

A. lusiocarpu. 383 Abrotme!la hnfllrlS V3f. apfrulata. 53

abscisic acid (ARA), 35. 153, 155. 156.

159, 160-4. 167. 168. 169

AOutilan thtophranil, 7 Ac(lOIl, 46. 187,212

It trumgium, 298

It omria. 26 A. papynXaTJkl. as p:l1l'h formers,

67.319

A tortiUis. 328 A. wrlidllllla. 36

AcanlhaH·:u~. 318 ACllula5poru. and racilitating

restoration. 366 acdimation. 1116 ,\elT. 7. 39, 198,201.204,206

A platanoidts. 297

11. pseudoplutunus. 42 A. runrum. 201. 257. 305

It sdCmarlnum. 257

A 5

INDEX

Arnbldopsis. 10. 124, 153. 155. 157,

'" A !haIUlna, 46. SO, 151. 304 and phytohormonl's. 168

mutants and phytohormone function, 168

Arubis hirsu/a, 208 AI

Cabombacl:ae, 135 Campwlhl'(a

IND EX

Cvrdyline austmJis. 19

Coriandrum saliwm. 151 Cornaceae. 52. 151. 27H. 298

Comus. 298

C. sanguinea. 278

Corokia macrocarpa. 52

Curtadma jubutu. 300 t·olylcdon. 18, 20

cotylcdonary axis

adjacent. 20

remote, 20

cotyledons. 8. 24-5. 126

buriaL 25

food storing. 25

function. 127 functional mor